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Haywardina Aczél (Diptera: Tephritidae)

Recognition
Species of Haywardina may be distinguished from other Tephritidae by the following combination of characters:     .
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Classification and Evolutionary Relationships
Order: Diptera. Family: Tephritidae. Subfamily: Trypetinae. Tribe: Carpomyini. Genus: Haywardina. Author: Aczél.
Haywardina belongs to the tribe Carpomyini, but its exact relationships within this group are not fully understood. It may be most closely related to Cryptodacus, Rhagoletotrypeta, and Zonosemata. The phylogenetic relationships among the species of Haywardina were analyzed by Norrbom (1994). Click here for a fuller discussion of Haywardina phylogeny.

Diversity & Distribution
Haywardina is endemic to the American tropics. There are 4 currently recognized species, known from Ecuador, Peru, and Argentina.
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Click here for list of included species and links to species pages.
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Names Used for this Genus
Haywardina Aczél 1951: 258, Type species: Tomoplagia cuculi Hendel, by original designation
Cryptoplagia Aczél 1951: 265, Type species: C. cuculiformis Aczél, by original designation. Synonymy (Norrbom 1994).
Click here to link to fly names database for full nomenclatural data.

Biology
The biology of Haywardina species is poorly known. A host plant is known only for H. cuculi Hendel, which has been reared from fruits of Solanum trichoneuron Lillo (Solanaceae) (Norrbom 1994). Numerous adults of that species, some in copulation, were collected on S. argentinum Bitter & Lillo, which also may be a host (Aczél 1951). Haywardina cuculiformis (Aczél) was also reared from fruits of an unidentified plant.

Economic Significance
Species of Haywardina are not considered economically important.

Comments
Norrbom (1994) revised and keyed the species of Haywardina and also proposed the synonymy of Cryptoplagia with Haywardina. Aczél (1951) probably proposed Haywardina and Cryptoplagia as separate monotypic genera because H. cuculi has several distinctive autapomorphies, but H. cuculiformis appears to be the species most closely related to it (see Haywardina phylogeny). By the generic limits proposed by Aczél, Cryptoplagia would include H. bimaculata (and perhaps also H. obscura), but it would be a paraphyletic taxon without also including H. cuculi. Rather than placing each of these three species in separate monotypic genera, Norrbom (1994) considered Haywardina and Cryptoplagia subjective synonyms. These names were published simultaneously by Aczél, but as recommended by the International Code of Zoological Nomenclature, Haywardina was chosen as the valid name because it was described prior to Cryptoplagia in Aczél's paper.

Haywardina bimaculata and cuculiformis each share a different apomorphy with Zonosemata, interpreted by Norrbom (1994) as homoplasy because of other character state distributions. It is possible that one of these species is a plesiomorphic member of Zonosemata, but, based on the characters analyzed, that hypothesis is less parsimonious than the hypothesis that Haywardina is monophyletic (see Haywardina phylogeny). The inclusion of H. obscura in Haywardina is tentative, because its relationships are less certain than those of the other species. Discovery of the female and study of its genitalia may clarify its relationships.

References
Click here to access fruit fly literature database
Aczél, M. L. 1951. Generos y especies neotropicales de la tribus "Trypetini". II. Dos generos y una especie nuevos. Acta Zool. Lilloana 12:253-278. [p. 265, description of Cryptoplagia, p. 265, description of Haywardina]
Foote, R. H. 1967. Family Tephritidae (Trypetidae, Trupaneidae). In N. Papavero, ed., A Catalogue of the Diptera of the Americas South of the United States. Departmento de Zoologia, Secretaria da Agricultura, São Paulo. Fasc. 57, 91 pp. [p. 21, 25, catalog]
Foote, R. H. 1980. Fruit fly genera south of the United States. United States Department of Agriculture, Technical Bulletin No. 1600, 79 pp. [p. 24, 31; review]
Norrbom, A. L. 1989. The status of Urophora acuticornis and U. sabroskyi (Diptera: Tephritidae). Entomol. News. 100: 59-66. [relationships, classification]
Norrbom, A. L. 1994. New species and phylogenetic analysis of Cryptodacus, Haywardina, and Rhagoletotrypeta (Diptera: Tephritidae). Insecta Mundi 8: 37-65. [revision, phylogenetic analysis]
Norrbom, A. L., L. E. Carroll, F. C. Thompson, I. M. White & A. Freidberg. 1999. Systematic database of names, pp. 65-251. In F. C. Thompson (ed.), Fruit Fly Expert Identification System and Systematic Information Database. Myia (1998) 9, vii + 524 pp. & Diptera Data Dissemination Disk (CD-ROM) (1998) 1. [p.   , catalog]

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